Homo sapiens · seed P00918 · 260 aa · family defined as ≥30% identity to that seed · compiled 09 August 2026
Every figure here is counted over the whole family rather than quoted from one entry.
232 distinct constructs across 1,490 entries. 771 polymer entities differ from the UniProt canonical sequence in some way, 40 carry a recognised expression tag and 0 carry a fusion partner.
"Differs from canonical" is not the same as "engineered". The canonical sequence is the full gene product, so a secreted protein whose structures all start after its signal peptide counts every one of them as different: lysozyme's most-used construct, residues 19–147 on 1,239 entities, is simply the mature protein. Read the construct column below for what was actually done, rather than this count.
| Entities | Length | Best (Å) | Best entry | What was made |
|---|---|---|---|---|
| 566 | 260 | 0.90 | 3K34 | matches the canonical sequence |
| 119 | 259 | 0.95 | 1LUG | residues 2-260 |
| 58 | 257 | 1.10 | 5NXW | residues 4-260 |
| 54 | 265 | 0.93 | 6ROB | matches the canonical sequence |
| 45 | 260 | 0.95 | 6YZN | A65S, N67Q, E69T +4 more |
| 44 | 257 | 1.12 | 6OTK | residues 4-260; A65S, N67Q, E69T +4 more |
| 39 | 258 | 1.00 | 4Q78 | residues 3-260 |
| 32 | 261 | 1.24 | 7Q0D | matches the canonical sequence |
| 29 | 263 | 1.12 | 9F2O | residues 29-291; G29M |
| 22 | 262 | 1.05 | 30TA | matches the canonical sequence |
| 15 | 263 | 1.06 | 5OGJ | matches the canonical sequence |
| 14 | 260 | 1.55 | 2FOY | residues 2-261 |
| 12 | 257 | 1.75 | 6G9U | residues 135-391; D136P, C174S |
| 12 | 259 | 1.80 | 1I9Q | residues 2-260; F130V |
| 12 | 260 | 1.04 | 7U5X | S2A |
| 11 | 260 | 1.07 | 4Q08 | A65S, N67K, I91T +3 more |
| 11 | 274 | 1.10 | 8Q3U | His6; C185S, C219S |
| 9 | 260 | 1.45 | 3OKV | A65S, N67Q |
| 9 | 263 | 1.19 | 9R0U | residues 29-291; G29A |
| 9 | 266 | 1.72 | 3FW3 | residues 19-284 |
| 8 | 260 | 1.05 | 1MOO | H64A |
| 8 | 260 | 1.54 | 2CBA | matches the canonical sequence |
| 7 | 257 | 1.98 | 7POM | residues 135-391; D136P, C174S, N346Q |
| 7 | 260 | 1.14 | 8RNS | A65S, N67K, E69N +6 more |
| 6 | 243 | 2.15 | 8DYQ | His6; 1 internal deletion; M1V, S23G, A24S +2 more |
Showing the 25 most-used of 232.
Columns where the wild-type residue still dominates but a real minority carries something else, which is a different question from "what varies across species".
| Oligomeric state | Chains | Entries | Share |
|---|---|---|---|
| monomeric | 1 | 1,376 | 92.3% |
| dimeric | 2 | 92 | 6.2% |
| tetrameric | 4 | 11 | 0.7% |
| trimeric | 3 | 8 | 0.5% |
| hexameric | 6 | 2 | 0.1% |
| pentameric | 5 | 1 | 0.1% |
911 entries have the depositor's assembly corroborated by PISA, 538 carry the depositor's word alone and 41 were assigned by PISA where the depositor gave none. The middle figure is not a disagreement: PISA may have returned nothing or never run. 5 entries carry more than one assembly with different chain counts, so they have no single answer to quote: 1Y7W, 3L14, 4LP6, 4Q0L, 5KU6.
| Domain | Source | Span (seed) | Chains |
|---|---|---|---|
| Alpha carbonic anhydrase | CATH 3.10.200.10 | 4–260 | 864 |
| Carbonic anhydrase | SCOP2B 8036258 | 4–260 | 1,149 |
| Carbonic anhydrase | SCOP2B 8034342 | 7–260 | 39 |
| Carbonic anhydrase | SCOP2B 8042916 | 9–260 | 42 |
| Carbonic anhydrase | SCOP2B 8065103 | 10–260 | 31 |
| Carbonic anhydrase | SCOP2B 8065111 | 10–260 | 18 |
| Component | Class | Name | Entries | Best (Å) |
|---|---|---|---|---|
| ZN | cofactor | Zinc Ion | 1,394 | 0.90 |
| GOL | cryoprotectant | Glycerol | 427 | 0.90 |
| DMS | cryoprotectant | Dimethyl Sulfoxide | 237 | 0.94 |
| HG | ion | Mercury (Ii) Ion | 154 | 0.93 |
| EDO | cryoprotectant | 1,2-Ethanediol | 84 | 0.99 |
| MBO | ligand | Mercuribenzoic Acid | 82 | 0.95 |
| SO4 | ion | Sulfate Ion | 79 | 1.20 |
| NA | ion | Sodium Ion | 59 | 0.94 |
| CO2 | solvent | Carbon Dioxide | 52 | 0.90 |
| BE7 | ligand | (4-Carboxyphenyl)(Chloro)mercury | 51 | 0.93 |
| BCN | buffer | Bicine | 45 | 0.98 |
| BCT | buffer | Bicarbonate Ion | 42 | 0.90 |
| CL | ion | Chloride Ion | 39 | 1.03 |
| ACT | cryoprotectant | Acetate Ion | 33 | 1.01 |
| AZM | ligand | 5-Acetamido-1,3,4-Thiadiazole-2-Sulfonamide | 31 | 1.10 |
| MES | buffer | 2-(N-Morpholino)-Ethanesulfonic Acid | 30 | 1.06 |
| TRS | buffer | 2-Amino-2-Hydroxymethyl-Propane-1,3-Diol | 28 | 1.09 |
| PEG | cryoprotectant | Di(Hydroxyethyl)ether | 28 | 0.99 |
| BGC | ligand | Beta-D-Glucopyranose | 27 | 0.93 |
| NI | ion | Nickel (Ii) Ion | 26 | 1.20 |
Parsed from the free text 1,369 depositors typed into
_exptl_crystal_grow.pdbx_details, out of 1,388
entries that recorded anything at all.
Median pH 7.8
(range 4.0 to 11.0).
1,490 entries carry a wwPDB validation report: 1,170 clean, 277 worth a check and 43 with something to explain. Median clashscore 3.9, median RSRZ outliers 1.94%, median R-free minus R-work 0.031. 1,359 have released structure factors.
| Organism | Entries | Best (Å) | Ligand-bound | Seed covered |
|---|---|---|---|---|
| Homo sapiens | 1,423 | 0.90 | 1341 | 100% |
| Bos taurus | 14 | 1.23 | 14 | 100% |
| Schistosoma mansoni | 10 | 1.60 | 10 | 92% |
| Mus musculus | 9 | 1.88 | 9 | 99% |
| Neisseria gonorrhoeae | 8 | 1.90 | 8 | 97% |
| Orthopoxvirus vaccinia | 5 | 1.42 | 0 | 85% |
| Unknown | 5 | 1.75 | 5 | 100% |
| Persephonella marina EX-H1 | 3 | 2.00 | 3 | 96% |
| Thermovibrio ammonificans | 2 | 1.69 | 2 | 97% |
| Chlamydomonas reinhardtii | 2 | 2.60 | 2 | 96% |
| Photobacterium profundum SS9 | 1 | 1.50 | 1 | 96% |
| Thermovibrio ammonificans HB-1 | 1 | 1.55 | 1 | 97% |
260 residues. Every identity figure in this document is measured against this sequence.
One record per paper, not per entry.
| Year | Citation |
|---|---|
| 2026 | 1-Aryl-6,7-Dimethoxy-3,4-Dihydroisoquinoline-2(1H)-Sulfonamides as hCA XII Selective Inhibitors: Experimental and Theoretical Studies to Interrogate the Isoform Selectivity. Chemmedchem doi:10.1002/cmdc.70390 |
| 2026 | X-ray analysis of complexes of carbonic anhydrase II with 1,3-oxazole-containing sulfonamide derivatives elucidates the structural basis for their exceptionally high inhibitory potency. Acta Crystallogr.,Sect.F doi:10.1107/S2053230X26002128 |
| 2026 | Discovery of a Mixed and Prodrug-Like Inhibition Mechanism for Phosphocoumarins and Phosphoquinolinones against Human Carbonic Anhydrases. J.Med.Chem. doi:10.1021/acs.jmedchem.6c00915 |
| 2026 | Achieving femtomolar affinities in structure-based drug design. Eur.Biophys.J. doi:10.1007/s00249-025-01812-5 |
| 2025 | X-ray crystallographic and kinetic studies of biguanide containing aryl sulfonamides as carbonic anhydrase inhibitors. Rsc Med Chem doi:10.1039/d4md01018c |
| 2025 | Design of Rigid Compounds to Enhance Selectivity for Carbonic Anhydrase IX. Chemistry doi:10.1002/chem.202404409 |
| 2025 | Di- meta -Substituted Fluorinated Benzenesulfonamides as Potent and Selective Anticancer Inhibitors of Carbonic Anhydrase IX and XII. J.Med.Chem. doi:10.1021/acs.jmedchem.5c01142 |
| 2025 | Affinity and Selectivity of Protein-Ligand Recognition: A Minor Chemical Modification Changes Carbonic Anhydrase Binding Profile. J.Med.Chem. doi:10.1021/acs.jmedchem.5c01421 |
| 2025 | Fast product release requires active-site water dynamics in carbonic anhydrase. Nat Commun doi:10.1038/s41467-025-59645-x |
| 2025 | Depsides from Origanum dictamnus and Satureja pilosa as selective inhibitors of carbonic anhydrases: Isolation, structure elucidation, X-ray crystallography. Arch Pharm doi:10.1002/ardp.202400823 |
| 2025 | An ureido-substituted benzenesulfonamide carbonic anhydrase inhibitor exerts a potent antitumor effect in vitro and in vivo. Exp Hematol Oncol doi:10.1186/s40164-025-00690-z |
| 2025 | Conformational flexibility of His200 enables catalytic activity in the T200H mutant of carbonic anhydrase II. Mol.Cells doi:10.1016/j.mocell.2025.100226 |
| 2025 | Dual inhibition of carbonic anhydrase IX and glutathione peroxidase 4 as a novel strategy for ferroptosis-induced tumor cell death. Eur.J.Med.Chem. doi:10.1016/j.ejmech.2025.118107 |
| 2025 | Structural Studies of the Dopamine D 4 Receptor Antagonist Sonepiprazole as an Inhibitor of Human Carbonic Anhydrases. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.5c00034 |
| 2025 | Unprecedented carbonic anhydrase inhibition mechanism: Targeting histidine 64 side chain through a halogen bond. Arch Pharm doi:10.1002/ardp.202400776 |
| 2025 | Tetrazole Is a Novel Zinc Binder Chemotype for Carbonic Anhydrase Inhibition. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.4c00562 |
| 2025 | Off-target binding of the histone deacetylase inhibitor vorinostat to carbonic anhydrase II and IX. Acta Crystallogr.,Sect.F doi:10.1107/S2053230X25007447 |
| 2025 | Targeting Human Carbonic Anhydrases with Novel Piperazine and Homopiperazine Benzenesulfonamides to Alleviate Paclitaxel-Induced Peripheral Neuropathy. J.Med.Chem. doi:10.1021/acs.jmedchem.5c02626 |
| 2025 | O-derivatization of natural tropolone and beta-thujaplicin leading to effective inhibitors of human carbonic anhydrases IX and XII. Eur.J.Med.Chem. doi:10.1016/j.ejmech.2025.117552 |
| 2025 | Protein crystallization and structure determination at room temperature in the CrystalChip. Febs Open Bio doi:10.1002/2211-5463.13932 |
| 2025 | Design, anticancer activity, and mechanistic evaluation of a novel class of selective human carbonic anhydrase IX inhibitors featuring a trifluorodihydroxypropanone pharmacophore. Eur.J.Med.Chem. doi:10.1016/j.ejmech.2025.118043 |
| 2025 | Structure-based design of an aromatic helical foldamer-protein interface. Chem Sci doi:10.1039/d5sc01826a |
| 2024 | Lasamide, a Potent Human Carbonic Anhydrase Inhibitor from the Market: Inhibition Profiling and Crystallographic Studies. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.4c00341 |
| 2024 | Exploring the binding mode of phenyl and vinyl boronic acids to human carbonic anhydrases. Int.J.Biol.Macromol. doi:10.1016/j.ijbiomac.2024.136873 |
| 2024 | Novel Carbonic Anhydrase Inhibitors with Dual-Tail Core Sulfonamide Show Potent and Lasting Effects for Glaucoma Therapy. J.Med.Chem. doi:10.1021/acs.jmedchem.3c02254 |
| 2024 | From X-ray crystallographic structure to intrinsic thermodynamics of protein-ligand binding using carbonic anhydrase isozymes as a model system. Iucrj doi:10.1107/S2052252524004627 |
| 2024 | Inhibition of Pseudomonas aeruginosa Carbonic Anhydrases, Exploring Ciprofloxacin Functionalization Toward New Antibacterial Agents: An In-Depth Multidisciplinary Study. J.Med.Chem. doi:10.1021/acs.jmedchem.4c01555 |
| 2024 | Study of Chalcogen Aspirin Derivatives with Carbonic Anhydrase Inhibitory Properties for Treating Inflammatory Pain. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.4c00284 |
| 2024 | Sulfonamide-incorporated bis( alpha-aminophosphonates) as promising carbonic anhydrase inhibitors: Design, synthesis, biological evaluation, and X-ray crystallographic studies. Arch Pharm doi:10.1002/ardp.202400038 |
| 2024 | XFEL structure of carbonic anhydrase II: a comparative study of XFEL, NMR, X-ray and neutron structures. Acta Crystallogr D Struct Biol doi:10.1107/S2059798324000482 |
| 2024 | Controlling the incorporation of fluorinated amino acids in human cells and its structural impact. Protein Sci. doi:10.1002/pro.4910 |
| 2024 | Benzoxaborinine, New Chemotype for Carbonic Anhydrase Inhibition: Ex Novo Synthesis, Crystallography, In Silico Studies, and Anti-Melanoma Cell Line Activity. J.Med.Chem. doi:10.1021/acs.jmedchem.4c01516 |
| 2024 | The dopamine D 2 receptors antagonist Veralipride inhibits carbonic anhydrases: solution and crystallographic insights on human isoforms. Chem Asian J doi:10.1002/asia.202400067 |
| 2024 | Targeted anticancer pre-vinylsulfone covalent inhibitors of carbonic anhydrase IX. Elife doi:10.7554/eLife.101401 |
| 2024 | Exploring the Polypharmacological Potential of PCI-27483: A Selective Inhibitor of Carbonic Anhydrases IX and XII. Acs Med.Chem.Lett. doi:10.1021/acsmedchemlett.4c00443 |
| 2024 | Machine Learning-Enhanced Quantum Chemistry-Assisted Refinement of the Active Site Structure of Metalloproteins. Inorg.Chem. doi:10.1021/acs.inorgchem.4c01274 |
| 2024 | 4-(Pyrazolyl)benzenesulfonamide Ureas as Carbonic Anhydrases Inhibitors and Hypoxia-Mediated Chemo-Sensitizing Agents in Colorectal Cancer Cells. J.Med.Chem. doi:10.1021/acs.jmedchem.4c01894 |
| 2024 | Microsecond Timescale Conformational Dynamics of a Small-Molecule Ligand within the Active Site of a Protein. Angew.Chem.Int.Ed.Engl. doi:10.1002/anie.202313947 |
| 2024 | Thia- and Seleno-Michael Reactions for the Synthesis of Carbonic Anhydrases Inhibitors. Chemmedchem doi:10.1002/cmdc.202400345 |
| 2024 | Directed Evolution of an Artificial Hydroxylase Based on a Thermostable Human Carbonic Anhydrase Protein Acs Catalysis doi:10.1021/acscatal.4c04163 |